鸭致病性E.coli外膜蛋白及相关毒力基因和新血清型发现及病原特性研究
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摘要
大肠杆菌(Escherichia coli,E.coli)是正常肠道菌群的组成部分,一些特殊血清型的E.coli对人和动物有病原性,动物的E.coli常常通过食物链的途径对人类健康造成严重的危害。随着集约化养殖的普及,鸭致病性E.coli对养鸭业的危害和导致的经济损失日益严重,已成为目前危害养鸭业最严重的传染病之一。现有禽病原性大肠杆菌(Avian Pathogenic Escherichia coli,APEC)的研究资料绝大多数来源于鸡,到目前为止,缺乏鸭源E.coli病原特性的系统研究。本研究旨在对我国西南地区规模化养鸭场鸭大肠杆菌病的致病性E.coli进行分离和血清型鉴定、了解鸭源致病性E.col外膜蛋白型(outer membrane protein patterns,OMP型)特征和相关毒力基因携带情况,对发现的一株致鸭腿水肿为特征的O46分离株的病原特性进行了系统研究,为阐明鸭源致病性E.col的致病机理和该病的防制提供科学数据。
1.规模化养鸭场鸭源致病性大肠杆菌的分离鉴定及病原特性和系统发育学分析
从我国四川、重庆、云南3个省(市)规模化养鸭场患典型大肠杆菌败血症雏鸭分离的282株致病性E.coli(临床分离株),通过玻板凝集和试管凝集试验,测定出210个分离株的血清型,占分离株的74.45%,分离株覆盖了37个血清型,其中O93、O78、O92、O76占43.8%(96/210)为优势血清型,O46、O32&O93混合型、O60&O93混合型为首次从鸭群中分离到。35株临床健康雏鸭泄殖腔分离的E.coli(泄殖腔分离株)鉴定出28株(包含14种血清型),其中O109和O154占46.4%为优势血清型。
对207个临床分离株进行了药敏试验,在所选的29种抗生素中,鸭大肠杆菌?分离株耐药性非常严重,平均耐药率28.83%,以多重耐药为主,耐10～18种药物的菌株占59.4%。
选取82个分离株(37个血清型,其中优势血清型各选8株共32株,其他血清型50株)进行致病性试验,以每株0.2ml(10~9CUF/ml)腿部肌肉接种7日龄鸭,确定出高致病株、中等致病株和低致病株分别占受试菌株的80.5%(66/82)、17.1%(14/82)和2.4%(2/82),四个优势血清型均为高、中致病株分别占受试菌株的84.4%(27/32)、15.6%(5/32)。
根据16SrRNA基因的序列同源性,对14个鸭致病性E.coli代表株进行系统发育分析,结果显示:14个代表株形成2个主要分枝,3株鸭致病性E.coli单独形成一个较远的分支:11株鸭致病性E.coli与人及其他动物源E.coli形成一个大的分支,6株与人的O157亲缘关系近,其中有2株与O157sakai聚为一支;另外5株与猪、牛、禽和人源E.coliK12亲缘关系近,同时还发现O血清型与菌株的亲缘关系无直接联系。
2.鸭源致病性大肠杆菌外膜蛋白型
测定了130株(34个血清型)鸭E.coli临床分离株外膜蛋型(OMP型),其中O93等优势血清型53株,O46等30个其他血清型77株,这些分离株共产生了4个OMP型,其中OMP-1型81株,占62.3%(81/130),覆盖O93、O92、O78和O76等29个血清型,为主要OMP型;根据GenBank中大肠杆菌K-12外膜蛋白A基因(ompA)的核苷酸序列设计引物,对9株优势血清型和1株O46血清型分离株的ompA进行克隆、序列测定和分析,发现优势血清型9个菌株的ompA开放性阅读框(ORF)长1053bp,编码由350个氨基酸组成的前外膜蛋白A(pro-OmpA),前21个氨基酸残基组成信号肽,成熟的OmpA由329个氨基酸残基组成;O46分离株的开放性阅读框全长1041bp,在400bp～411bp的地方缺失了12个碱基,成熟的OmpA由325个氨基酸残基组成;10个分离株的ompA序列其核苷酸序列高度同源,达95.8%～100%。对10个分离株ompA进行系统发育分析,O46分离株单独聚为一支,其他9个分离株聚为一支;菌株血清型、外膜蛋白型与ompA的亲缘关系无直接联系。
3.鸭源致病性大肠杆菌相关毒力基因分析
应用PCR结合核酸序列测定对210株临床分离株和28株泄殖腔分离株进行包括强毒力岛(HPI)中的鼠疫菌素受体基因(fyuA)和铁调节蛋基因(irp2)、I型菌毛必须蛋白基因(fimC)、P型菌毛结构基因(papA)和血清耐受基因(iss)进行检测,结果表明:fyuA、irp2、fimC、papA和iss在临床分离株的携带率分别为41.0%、43.3%、92.9%、97.6%和96.7%,在泄殖腔分离株的携带率分别为21.4%、25.0%、92.9%、100%和92.9%,临床分离株和泄殖腔分离株Iss、I型菌毛和P型菌毛基因携带率差异不显著(P＞0.05),但P型菌毛基因的携带率均显著高于其他宿主(鸡、猪和人)源E.coli;HPI毒力岛在鸭源E.coli中分布较广,其携带率表现为临床分离株极显著高于泄殖腔分离株。HPI毒力岛的携带率与菌株的致病性呈明显的正相关,与O78等特定的血清型有一定的关系。临床分离株有37.6%(79/210)同时携带fyuA、irp2、fimC、iss和papA基因,极显著高于泄殖腔分离株14.3%(4/28)(P＜0.01)。
4.鸭源致病性大肠杆菌志贺毒素相关基因分子流行病学
根据GenBank中编码大肠杆菌stx1(志贺毒素1基因)、stx2(志贺毒素2基因)、hlyA(溶血素基因)和eaeA(黏附素基因)的特异性序列,设计合成4对引物,建立了快速鉴定产志贺毒素大肠杆菌(STEC)多重PCR方法。该方法能直接特异性鉴定多种毒力因子,检测敏感度细菌纯培养物为10~3CFU/ml。用此方法结合核酸序列测定对210株临床分离株和28株泄殖腔分离株进行相关基因检测,结果表明:临床分离株有6株具有six1序列、4株具有hlyA序列、1株具有eaeA序列;stx1、hlyA和eaeA阳性率分别为2.85%(6/210)、1.90%(4/210)和0.47%(1/210);5株基因型为stx1~+,3株基因型为hlyA~+,1株基因型为stx1~++hlyA~++eaeA~+,分别为2.38%(5/210)、1.43%(3/210)和0.47%(1/210);临床分离株stx2均为阴性。泄殖腔分离株未检出这4种基因。对所有阳性基因进行序列测定及分析,hlyA与GenBank中提供的EHECO157:H7 Sakai株序列的同源性为100%,stx1和eaeA与O157:H7 Sakai株序列高度同源。在发病鸭中发现并分离到O158、O60、O78、O36、O77、O46、O137和O192(携带stx1、hlyA、eaeA)的非-O157的STEC菌株,具有重要的公共卫生学意义。
5.一株致鸭腿水肿为特征的O46大肠杆菌的发现及其实验病理模型建立
从临床急性死亡,以腿水肿为特征的病鸭群中分离的SYW004株,通过形态学、培养特性、生化特征的研究和血清型鉴定,确定为大肠杆菌O46血清型;16S rRNA基因序列分析表明,该分离株与人源志贺氏菌及大肠杆菌O157:H7的亲缘关系最近,序列同源性为99%～99.5%,而和鸡源E.coli亲缘关系较远;对hlyA等9种毒力基因检测表明,O46分离株携带hlyA、fyuA、irp2、fimC、papA和泌六种毒力基因,是一株携带多种毒力基因的出血性大肠杆菌(EHEC);该分离株对雏鸭具有高致病性,实验病理模型能复制出与自然感染相同的病例,组织病理学检查表明,肝脏、肺脏、心脏、肾脏、肠道和脑等器官出现不同程度的组织细胞变性、坏死和大面积淤血、出血;超薄切片电镜观察可在心脏、肺脏、脾脏、肝脏和小肠中观察到大肠杆菌,其侵害的主要靶细胞包括淋巴细胞、巨噬细胞、肠道上皮细胞、心肌纤维细胞。超微结构变化最早发生于心脏、肝脏和肠道,各种细胞的变化主要为肿胀、坏死,染色质凝聚,线粒体肿胀,嵴断裂、扭曲变形,心肌肌原纤维间距增宽、溶解断裂,肠腺上皮细胞肿胀,微绒毛脱落、固有层充血出血。
6.鸭源致病性大肠杆菌O46分离株对人工感染鸭的侵染规律
为了观察O46分离株在人工感染鸭体内的分布,以O46定型血清建立免疫组化方法,应用建立的方法对O46分离株人工感染雏鸭的不同时间的不同组织器官进行观察。结果显示:免疫组化的方法只对大肠杆菌O46感染致死鸭的肝脏、脾脏、肾脏呈现阳性反应而与大肠杆菌O78、鸭瘟、鸭源禽流感、鸭疫里默氏杆菌、沙门氏菌、多杀性巴氏杆菌感染致死鸭的肝组织呈现阴性反应,具有直观、敏感、特异性强的优点,能够进行精确抗原定位;实验结果还表明,O46分离株能在鸭体内进行大面积的侵嗜,心脏、肺脏、脾脏、肾脏和肠道是感染的主要靶器官,阳性信号最早出现于心脏,气管未检测到细菌抗原。我们的实验数据可以为大肠杆菌O46分离株在体内分布和复制提供重要的证据,同时可以为O46分离株感染的致病机制提供一定的知识。
Escherichia coli(E.coli)are a common member of the intestinal normal flora,but some E.coli strains belonging to certain serotypes are pathogenic to human and animals. Pathogenic E.coli of animals may pose a serious threat to human health throughout the food chain.With the popularization of intensive raising duck,pathogenic E.coli is becoming an increasingly serious problem because of great economic losses in duck industry.Unfortunately,most of the researches concerning avian pathogenic Escherichia coli(APEC)were focused on E.coli originated from chickens,and only have a few of pathogenic characteristics of duck-origin E.coli been reported in the past years.The objectives of this study were to isolate pathogenic E.coli strains from different large-scale duck farms in southwest China,and to investigate their serotypes,outer membrane protein patterns(OMP)and virulence-related genes.A novel serotype O46 strain,which cause disease characterized by swollen leg,was isolated and its biological features were studied intensively in this study.The results of this research will provide a scientific data for clarifying the pathogenesis,and for controlling and preventing the diseases caused by these E coli.
1.Isolation,identification and phylogenetic analysis of duck pathogenic E.coli from large-scale duck farms.
Two hundreds and eighty two strains of pathogenic E.coli were isolated from septicemia ducks of large-scale duck farms in Sichuan,Chongqing and Yunan provinces in China.Two hundreds and ten out of the 282 isolates(74.45%)were determined for serotypes by using slide agglutination test and tube agglutination test.The results showed that the isolates were related to thirty seven serotypes.Serotypes O93,O78,O92 and O76 were predominant serotypes,accounting for 43.8%(96/210),and serotype O46,O32&O93 and serotype O60&O93 were mixed-serotype which were isolated from ducks for the first time.
Twenty eight strains belonging to 14 serotypes were identified from 35 isolates from cloaca of clinically healthy ducklings,and O109 and O154,46.4%of the total,were predominant serotypes.
Antibiotic sensitivity tests were performed for 207 isolates and these isolates showed serious antibiotic resistance to 29 selected antibiotics.The average antibiotic resistance rate was 28.83%while most of strains showed multi-resistance and 59.4%of swains were resistant against 10 to 18 antibiotics.
Eighty two isolates belonging to 37 serotypes(8 strains from each prominent serotype and 50 strains from other serotypes)were selected and their pathogenicity was evaluated after injection into leg muscle of 7-day ducklings(0.2ml of 10~9CUF/ml each).The results showed that high-pathogenicity strains,middle-pathogenicity strains and low-pathogenicity strains accounted for 80.5%(66/82),17.1%(14/82)and 2.4%(2/82)of total isolates tested,respectively.It was also found that all strains belonging to predominant serotypes were high- or middle-pathogenicity strains,accounting for 84.4%(27/32)or 15.6%(5/32) of tested strains,respectively.
Phylogenetic analysis was conducted according to the 16S rRNA gene sequences of E.coli from 14 typical pathogenic strains of duck and from other animals.The results demonstrated that phylogenetic tree divided into two clusters.The first cluster,having longer branch length,consisted of 3 duck-origin pathogenic strains while the second cluster included 11 duck-origin pathogenic strains and some strains from human and other animals.Among those 11 strains,6 strains had a close relationship with E.coli serotype O157 of human-origin(2 strains had a close relationship with O157 sakai strain),while the other 5 strains showed close relationship with E.coli isolates from pig,cattle,poultry,or E.coli K12 strain of human.It is also revealed that there is no direct correlation between O serotypes and phylogenitic relationship of strains.
2.Determination of the outer membrane protein(OMP)patterns of pathogenic E.coli isolated from ducks
The outer membrane protein patterns(OMP)of 130 pathogenic E.coli isolates belonging to 34 serotypes from ducks were determined in this study.Among these 130 strains,53 isolates belonged to prominent serotypes such as O93,while 77 strains were in other 35 serotypes such as O46.The results revealed 4 different OMP patterns in all strains tested.62.3%(81/130)of strains were identified to be OMP-1 with 29 serotypes,including O93,O92,O78 and O76,suggesting OMP-1 was the main OMP.Subsequently,a primer pair for amplification of ompA gene was also designed according to sequence of the E.coli K-12 available in Genbank.The ompA gene were then amplified and sequenced from 9 prominent serotype strains and 1 serotype O46 strain.Sequence analysis revealed that the ompA gene of all 9 prominent serotype strains open reading frame(ORF)with a size of 1053bp,which encoded a 350 amino acid protein,pro-OmpA.The first 21amino acids encoded a signal peptide and the mature OmpA protein was composed of 329 amino acids. The ORF of ompA gene from O46 serotype strain was 1041bp in length with 12bp depletion at the site of 400bp～411bp compared to other serotype strains,and the mature OmpA protein contained 325 amino acids.It was also found that the nucleotide sequences of ompA gene were highly homological with an identity ranging from 95.8 to 100%among 10 pathogenic E.coli strains from ducks.Phylogentic analysis of OmpA from 10 isolates indicated that O46 serotype form a single branch while other 9 strains consisted of the second branch.The study also indicated that there is no direct correlation between serotype, OMP and OmpA phylogenetic relationship
3.Analysis of virulence-related genes of duck pathogenic E.coli isolates
In this part of the study,pesticin receptor gene(fyuA)within the highly pathogenecity islands(HPI),iron regulatory protein gene(irp),type 1 pilus essential protein gene(fimC), the pap pilus structural protein gene(papA)and serum tolerance associated gene(iss)of the 210 pathogenic E.coli strains isolated from ducks showing clinical signs(clinical isolate) and the 28 E.coli strains isolated from clinically healthy ducklings by cloacal swabs(cloacal isolate)were determined by using polymerase chain reaction(PCR) combined with nucleic acid sequence analysis.The results showed that fyuA,irp2,fimC, papA and iss were detected in 41.0%,43.8%,92.9%,97.6%,and 96.7%of clinical isolates, and in 21.4%,25.0%,92.9%,100%and 92.9%of cloacal isolates,respectively. Statistically,there were no significant difference(P＞0.05)between clinical isolates and cloacal isolates for iss,fimC and papA gene detection.For papA gene detection,there were significant differences(P＜0.05)between the clinical isolates and isolates from other hosts such as chickens,swine and humans.The gene of fyuA was widely distributed in ducks and its detection rate in ducks showing clinical signs was significantly higher (P＜0.05)than that in clinically healthy duckling.There was a close positive correlation between the fyuA gene and the pathogenicity of E.coli.Furthermore,the gene of fyuA was closely related with the specific serotype of O78.The detection rate of fyuA,irp2,fimC,iss and papA2 in ducks showing clinical signs(37.6%,79/210)was significantly higher(P＜0.01)than that in clinically healthy ducklings(14.3%,4/28).
4.Molecular epidemiology of Shinga toxin genes in E.coli from ducks.
Four sets of primers were designed for stx1,stx2,hlyA and eaeA according to the sequences published in Genbank,and a multiplex PCR assay was established for rapid detection of shiga toxin producing E.coli.This PCR method was able to detect 4 virulence factors specifically and with a sensitivity of 10~3CFU/ml of pure bacterial culture.Two hundreds and ten of clinical isolates and 28 of cloacal isolates were detected by the multiplex PCR.The results showed that 6 strains of the 210 clinical isolates were stx1 positive,4 strains were hlyA positive and 1 strain was eaeA positive.The positive rate of stx1、hlyA and eaeA were 2.85%(6/210),1.90%(4/210)and 0.47%(1/210),respectively. Among the 210 clinical isolates studied,5 strains belonged to genotype stx1~+,3 strains belonged to genotype hlyA~+ and 1 strains belonged to genotype stx1~++hlyA~++eaeA~+, accounting for 2.38%(5/210),1.43%(3/210)and 0.47%(1/210),respectively.All clinical isolates were str2 negative while all cloacal isolates were negative for all interested genes in this study.Sequencing analysis of all positive PCR products found hlyA gene have 100% identity with that of EHEC O157:H7 Sakai strain in Genbank,while stx1 and eaeA have high homology with that of STEC.In this study,non-O157 STEC strains carrying stx1, hlyA and eaeA,and belonging to serotypes O158,O60,O78,O36,O77,O46,O137 and O192,were isolated from sick ducks.This inferred that E.coli strains from duck-origin may be an important problem in public health.
5.Identification of a serotype O46 strain from ducks characterized by swelling leg and establishment of experimental pathological model.
A duck-origin E.coli strain,SYW004,was isolated from a flock of sick ducks with clinical signs characterized by acute death and swollen leg,and proved to be serotype O46 after morphological investigation,biochemical study and serotype analysis.16S rRNA gene sequencing revealed that SYW004 strain had a close relationship to the human shigella and E.coli O157:H7(99～99.5%identity)but a far relationship with E.coli of chicken.
Detection of 9 virulence genes demonstrated that this isolate carded 6 virulencerelated genes including hlyA,fyuA,irp2,fimC,papa and iss,suggesting it is an EHEC carrying multiple virulence genes.
The SYW004 strain showed high pathogenicity in ducklings.Experimental model of infection closely resembled the natural infection.Autopsy,histopathologic and ultra-structural pathological observation showed that the major pathological features were characterized by wide fibrinous inflammation in the serosa.Hyperemia,hemorrhage and inflammatory cell infiltrate were observed in heart,liver,spleen,lung,kidney,bursa of Fabricius,small intestine,pancreas and brain.Degeneration,necrosis or apoptosis were found in renal tubular epithelial cells,liver cells,cardiac muscle cells and intestinal epithelial cells.Lymphocytes decreased in bursa of Fabriscius.E.coli was observed on ultra-thin sections of heart,lung,spleen,liver and small intestine under electron microscope.The target cells were lymphocytes,phagocytes,intestinal epithelial cells and cardiac fiber.
6.Infection pattern of E.coli O46 isolate in an experimental model in ducks
To investigate the distribution of the O46 strain isolated in this study,an immunohisto chemical assay was developed with O46-specific antiserum and used to detect the bacteria of different organs at different times after experimental infection with the O46 isolate.The results showed this assay have good specificity.Samples of liver,spleen and kidney from ducks infected with O46 isolate showed positive results by this assay,while liver samples from dead ducks infected with E.coli O78,duck riemenella nantepestifer,duck plague virus,duck-origin avian influenza virus,salmonella or pasteurella multocida showed negative.With the advantages of direct,sensitivity and specificity,this assay could be used for accurate antigen localization.The study also indicated that the O46 isolates have wide tropism while heart,lung,spleen,kidney and intestine were found to be the main target organs.The bacteria were detected first in heart while bacteria could not be detected in trachea.Those findings provided important information for the in vivo distribution and proliferation of the O46 strain and accumulated useful knowledge for pathogenesis of this isolate.
1.规模化养鸭场鸭源致病性大肠杆菌的分离鉴定及病原特性和系统发育学分析
从我国四川、重庆、云南3个省(市)规模化养鸭场患典型大肠杆菌败血症雏鸭分离的282株致病性E.coli(临床分离株),通过玻板凝集和试管凝集试验,测定出210个分离株的血清型,占分离株的74.45%,分离株覆盖了37个血清型,其中O93、O78、O92、O76占43.8%(96/210)为优势血清型,O46、O32&O93混合型、O60&O93混合型为首次从鸭群中分离到。35株临床健康雏鸭泄殖腔分离的E.coli(泄殖腔分离株)鉴定出28株(包含14种血清型),其中O109和O154占46.4%为优势血清型。
对207个临床分离株进行了药敏试验,在所选的29种抗生素中,鸭大肠杆菌?分离株耐药性非常严重,平均耐药率28.83%,以多重耐药为主,耐10～18种药物的菌株占59.4%。
选取82个分离株(37个血清型,其中优势血清型各选8株共32株,其他血清型50株)进行致病性试验,以每株0.2ml(10~9CUF/ml)腿部肌肉接种7日龄鸭,确定出高致病株、中等致病株和低致病株分别占受试菌株的80.5%(66/82)、17.1%(14/82)和2.4%(2/82),四个优势血清型均为高、中致病株分别占受试菌株的84.4%(27/32)、15.6%(5/32)。
根据16SrRNA基因的序列同源性,对14个鸭致病性E.coli代表株进行系统发育分析,结果显示:14个代表株形成2个主要分枝,3株鸭致病性E.coli单独形成一个较远的分支:11株鸭致病性E.coli与人及其他动物源E.coli形成一个大的分支,6株与人的O157亲缘关系近,其中有2株与O157sakai聚为一支;另外5株与猪、牛、禽和人源E.coliK12亲缘关系近,同时还发现O血清型与菌株的亲缘关系无直接联系。
2.鸭源致病性大肠杆菌外膜蛋白型
测定了130株(34个血清型)鸭E.coli临床分离株外膜蛋型(OMP型),其中O93等优势血清型53株,O46等30个其他血清型77株,这些分离株共产生了4个OMP型,其中OMP-1型81株,占62.3%(81/130),覆盖O93、O92、O78和O76等29个血清型,为主要OMP型;根据GenBank中大肠杆菌K-12外膜蛋白A基因(ompA)的核苷酸序列设计引物,对9株优势血清型和1株O46血清型分离株的ompA进行克隆、序列测定和分析,发现优势血清型9个菌株的ompA开放性阅读框(ORF)长1053bp,编码由350个氨基酸组成的前外膜蛋白A(pro-OmpA),前21个氨基酸残基组成信号肽,成熟的OmpA由329个氨基酸残基组成;O46分离株的开放性阅读框全长1041bp,在400bp～411bp的地方缺失了12个碱基,成熟的OmpA由325个氨基酸残基组成;10个分离株的ompA序列其核苷酸序列高度同源,达95.8%～100%。对10个分离株ompA进行系统发育分析,O46分离株单独聚为一支,其他9个分离株聚为一支;菌株血清型、外膜蛋白型与ompA的亲缘关系无直接联系。
3.鸭源致病性大肠杆菌相关毒力基因分析
应用PCR结合核酸序列测定对210株临床分离株和28株泄殖腔分离株进行包括强毒力岛(HPI)中的鼠疫菌素受体基因(fyuA)和铁调节蛋基因(irp2)、I型菌毛必须蛋白基因(fimC)、P型菌毛结构基因(papA)和血清耐受基因(iss)进行检测,结果表明:fyuA、irp2、fimC、papA和iss在临床分离株的携带率分别为41.0%、43.3%、92.9%、97.6%和96.7%,在泄殖腔分离株的携带率分别为21.4%、25.0%、92.9%、100%和92.9%,临床分离株和泄殖腔分离株Iss、I型菌毛和P型菌毛基因携带率差异不显著(P＞0.05),但P型菌毛基因的携带率均显著高于其他宿主(鸡、猪和人)源E.coli;HPI毒力岛在鸭源E.coli中分布较广,其携带率表现为临床分离株极显著高于泄殖腔分离株。HPI毒力岛的携带率与菌株的致病性呈明显的正相关,与O78等特定的血清型有一定的关系。临床分离株有37.6%(79/210)同时携带fyuA、irp2、fimC、iss和papA基因,极显著高于泄殖腔分离株14.3%(4/28)(P＜0.01)。
4.鸭源致病性大肠杆菌志贺毒素相关基因分子流行病学
根据GenBank中编码大肠杆菌stx1(志贺毒素1基因)、stx2(志贺毒素2基因)、hlyA(溶血素基因)和eaeA(黏附素基因)的特异性序列,设计合成4对引物,建立了快速鉴定产志贺毒素大肠杆菌(STEC)多重PCR方法。该方法能直接特异性鉴定多种毒力因子,检测敏感度细菌纯培养物为10~3CFU/ml。用此方法结合核酸序列测定对210株临床分离株和28株泄殖腔分离株进行相关基因检测,结果表明:临床分离株有6株具有six1序列、4株具有hlyA序列、1株具有eaeA序列;stx1、hlyA和eaeA阳性率分别为2.85%(6/210)、1.90%(4/210)和0.47%(1/210);5株基因型为stx1~+,3株基因型为hlyA~+,1株基因型为stx1~++hlyA~++eaeA~+,分别为2.38%(5/210)、1.43%(3/210)和0.47%(1/210);临床分离株stx2均为阴性。泄殖腔分离株未检出这4种基因。对所有阳性基因进行序列测定及分析,hlyA与GenBank中提供的EHECO157:H7 Sakai株序列的同源性为100%,stx1和eaeA与O157:H7 Sakai株序列高度同源。在发病鸭中发现并分离到O158、O60、O78、O36、O77、O46、O137和O192(携带stx1、hlyA、eaeA)的非-O157的STEC菌株,具有重要的公共卫生学意义。
5.一株致鸭腿水肿为特征的O46大肠杆菌的发现及其实验病理模型建立
从临床急性死亡,以腿水肿为特征的病鸭群中分离的SYW004株,通过形态学、培养特性、生化特征的研究和血清型鉴定,确定为大肠杆菌O46血清型;16S rRNA基因序列分析表明,该分离株与人源志贺氏菌及大肠杆菌O157:H7的亲缘关系最近,序列同源性为99%～99.5%,而和鸡源E.coli亲缘关系较远;对hlyA等9种毒力基因检测表明,O46分离株携带hlyA、fyuA、irp2、fimC、papA和泌六种毒力基因,是一株携带多种毒力基因的出血性大肠杆菌(EHEC);该分离株对雏鸭具有高致病性,实验病理模型能复制出与自然感染相同的病例,组织病理学检查表明,肝脏、肺脏、心脏、肾脏、肠道和脑等器官出现不同程度的组织细胞变性、坏死和大面积淤血、出血;超薄切片电镜观察可在心脏、肺脏、脾脏、肝脏和小肠中观察到大肠杆菌,其侵害的主要靶细胞包括淋巴细胞、巨噬细胞、肠道上皮细胞、心肌纤维细胞。超微结构变化最早发生于心脏、肝脏和肠道,各种细胞的变化主要为肿胀、坏死,染色质凝聚,线粒体肿胀,嵴断裂、扭曲变形,心肌肌原纤维间距增宽、溶解断裂,肠腺上皮细胞肿胀,微绒毛脱落、固有层充血出血。
6.鸭源致病性大肠杆菌O46分离株对人工感染鸭的侵染规律
为了观察O46分离株在人工感染鸭体内的分布,以O46定型血清建立免疫组化方法,应用建立的方法对O46分离株人工感染雏鸭的不同时间的不同组织器官进行观察。结果显示:免疫组化的方法只对大肠杆菌O46感染致死鸭的肝脏、脾脏、肾脏呈现阳性反应而与大肠杆菌O78、鸭瘟、鸭源禽流感、鸭疫里默氏杆菌、沙门氏菌、多杀性巴氏杆菌感染致死鸭的肝组织呈现阴性反应,具有直观、敏感、特异性强的优点,能够进行精确抗原定位;实验结果还表明,O46分离株能在鸭体内进行大面积的侵嗜,心脏、肺脏、脾脏、肾脏和肠道是感染的主要靶器官,阳性信号最早出现于心脏,气管未检测到细菌抗原。我们的实验数据可以为大肠杆菌O46分离株在体内分布和复制提供重要的证据,同时可以为O46分离株感染的致病机制提供一定的知识。
Escherichia coli(E.coli)are a common member of the intestinal normal flora,but some E.coli strains belonging to certain serotypes are pathogenic to human and animals. Pathogenic E.coli of animals may pose a serious threat to human health throughout the food chain.With the popularization of intensive raising duck,pathogenic E.coli is becoming an increasingly serious problem because of great economic losses in duck industry.Unfortunately,most of the researches concerning avian pathogenic Escherichia coli(APEC)were focused on E.coli originated from chickens,and only have a few of pathogenic characteristics of duck-origin E.coli been reported in the past years.The objectives of this study were to isolate pathogenic E.coli strains from different large-scale duck farms in southwest China,and to investigate their serotypes,outer membrane protein patterns(OMP)and virulence-related genes.A novel serotype O46 strain,which cause disease characterized by swollen leg,was isolated and its biological features were studied intensively in this study.The results of this research will provide a scientific data for clarifying the pathogenesis,and for controlling and preventing the diseases caused by these E coli.
1.Isolation,identification and phylogenetic analysis of duck pathogenic E.coli from large-scale duck farms.
Two hundreds and eighty two strains of pathogenic E.coli were isolated from septicemia ducks of large-scale duck farms in Sichuan,Chongqing and Yunan provinces in China.Two hundreds and ten out of the 282 isolates(74.45%)were determined for serotypes by using slide agglutination test and tube agglutination test.The results showed that the isolates were related to thirty seven serotypes.Serotypes O93,O78,O92 and O76 were predominant serotypes,accounting for 43.8%(96/210),and serotype O46,O32&O93 and serotype O60&O93 were mixed-serotype which were isolated from ducks for the first time.
Twenty eight strains belonging to 14 serotypes were identified from 35 isolates from cloaca of clinically healthy ducklings,and O109 and O154,46.4%of the total,were predominant serotypes.
Antibiotic sensitivity tests were performed for 207 isolates and these isolates showed serious antibiotic resistance to 29 selected antibiotics.The average antibiotic resistance rate was 28.83%while most of strains showed multi-resistance and 59.4%of swains were resistant against 10 to 18 antibiotics.
Eighty two isolates belonging to 37 serotypes(8 strains from each prominent serotype and 50 strains from other serotypes)were selected and their pathogenicity was evaluated after injection into leg muscle of 7-day ducklings(0.2ml of 10~9CUF/ml each).The results showed that high-pathogenicity strains,middle-pathogenicity strains and low-pathogenicity strains accounted for 80.5%(66/82),17.1%(14/82)and 2.4%(2/82)of total isolates tested,respectively.It was also found that all strains belonging to predominant serotypes were high- or middle-pathogenicity strains,accounting for 84.4%(27/32)or 15.6%(5/32) of tested strains,respectively.
Phylogenetic analysis was conducted according to the 16S rRNA gene sequences of E.coli from 14 typical pathogenic strains of duck and from other animals.The results demonstrated that phylogenetic tree divided into two clusters.The first cluster,having longer branch length,consisted of 3 duck-origin pathogenic strains while the second cluster included 11 duck-origin pathogenic strains and some strains from human and other animals.Among those 11 strains,6 strains had a close relationship with E.coli serotype O157 of human-origin(2 strains had a close relationship with O157 sakai strain),while the other 5 strains showed close relationship with E.coli isolates from pig,cattle,poultry,or E.coli K12 strain of human.It is also revealed that there is no direct correlation between O serotypes and phylogenitic relationship of strains.
2.Determination of the outer membrane protein(OMP)patterns of pathogenic E.coli isolated from ducks
The outer membrane protein patterns(OMP)of 130 pathogenic E.coli isolates belonging to 34 serotypes from ducks were determined in this study.Among these 130 strains,53 isolates belonged to prominent serotypes such as O93,while 77 strains were in other 35 serotypes such as O46.The results revealed 4 different OMP patterns in all strains tested.62.3%(81/130)of strains were identified to be OMP-1 with 29 serotypes,including O93,O92,O78 and O76,suggesting OMP-1 was the main OMP.Subsequently,a primer pair for amplification of ompA gene was also designed according to sequence of the E.coli K-12 available in Genbank.The ompA gene were then amplified and sequenced from 9 prominent serotype strains and 1 serotype O46 strain.Sequence analysis revealed that the ompA gene of all 9 prominent serotype strains open reading frame(ORF)with a size of 1053bp,which encoded a 350 amino acid protein,pro-OmpA.The first 21amino acids encoded a signal peptide and the mature OmpA protein was composed of 329 amino acids. The ORF of ompA gene from O46 serotype strain was 1041bp in length with 12bp depletion at the site of 400bp～411bp compared to other serotype strains,and the mature OmpA protein contained 325 amino acids.It was also found that the nucleotide sequences of ompA gene were highly homological with an identity ranging from 95.8 to 100%among 10 pathogenic E.coli strains from ducks.Phylogentic analysis of OmpA from 10 isolates indicated that O46 serotype form a single branch while other 9 strains consisted of the second branch.The study also indicated that there is no direct correlation between serotype, OMP and OmpA phylogenetic relationship
3.Analysis of virulence-related genes of duck pathogenic E.coli isolates
In this part of the study,pesticin receptor gene(fyuA)within the highly pathogenecity islands(HPI),iron regulatory protein gene(irp),type 1 pilus essential protein gene(fimC), the pap pilus structural protein gene(papA)and serum tolerance associated gene(iss)of the 210 pathogenic E.coli strains isolated from ducks showing clinical signs(clinical isolate) and the 28 E.coli strains isolated from clinically healthy ducklings by cloacal swabs(cloacal isolate)were determined by using polymerase chain reaction(PCR) combined with nucleic acid sequence analysis.The results showed that fyuA,irp2,fimC, papA and iss were detected in 41.0%,43.8%,92.9%,97.6%,and 96.7%of clinical isolates, and in 21.4%,25.0%,92.9%,100%and 92.9%of cloacal isolates,respectively. Statistically,there were no significant difference(P＞0.05)between clinical isolates and cloacal isolates for iss,fimC and papA gene detection.For papA gene detection,there were significant differences(P＜0.05)between the clinical isolates and isolates from other hosts such as chickens,swine and humans.The gene of fyuA was widely distributed in ducks and its detection rate in ducks showing clinical signs was significantly higher (P＜0.05)than that in clinically healthy duckling.There was a close positive correlation between the fyuA gene and the pathogenicity of E.coli.Furthermore,the gene of fyuA was closely related with the specific serotype of O78.The detection rate of fyuA,irp2,fimC,iss and papA2 in ducks showing clinical signs(37.6%,79/210)was significantly higher(P＜0.01)than that in clinically healthy ducklings(14.3%,4/28).
4.Molecular epidemiology of Shinga toxin genes in E.coli from ducks.
Four sets of primers were designed for stx1,stx2,hlyA and eaeA according to the sequences published in Genbank,and a multiplex PCR assay was established for rapid detection of shiga toxin producing E.coli.This PCR method was able to detect 4 virulence factors specifically and with a sensitivity of 10~3CFU/ml of pure bacterial culture.Two hundreds and ten of clinical isolates and 28 of cloacal isolates were detected by the multiplex PCR.The results showed that 6 strains of the 210 clinical isolates were stx1 positive,4 strains were hlyA positive and 1 strain was eaeA positive.The positive rate of stx1、hlyA and eaeA were 2.85%(6/210),1.90%(4/210)and 0.47%(1/210),respectively. Among the 210 clinical isolates studied,5 strains belonged to genotype stx1~+,3 strains belonged to genotype hlyA~+ and 1 strains belonged to genotype stx1~++hlyA~++eaeA~+, accounting for 2.38%(5/210),1.43%(3/210)and 0.47%(1/210),respectively.All clinical isolates were str2 negative while all cloacal isolates were negative for all interested genes in this study.Sequencing analysis of all positive PCR products found hlyA gene have 100% identity with that of EHEC O157:H7 Sakai strain in Genbank,while stx1 and eaeA have high homology with that of STEC.In this study,non-O157 STEC strains carrying stx1, hlyA and eaeA,and belonging to serotypes O158,O60,O78,O36,O77,O46,O137 and O192,were isolated from sick ducks.This inferred that E.coli strains from duck-origin may be an important problem in public health.
5.Identification of a serotype O46 strain from ducks characterized by swelling leg and establishment of experimental pathological model.
A duck-origin E.coli strain,SYW004,was isolated from a flock of sick ducks with clinical signs characterized by acute death and swollen leg,and proved to be serotype O46 after morphological investigation,biochemical study and serotype analysis.16S rRNA gene sequencing revealed that SYW004 strain had a close relationship to the human shigella and E.coli O157:H7(99～99.5%identity)but a far relationship with E.coli of chicken.
Detection of 9 virulence genes demonstrated that this isolate carded 6 virulencerelated genes including hlyA,fyuA,irp2,fimC,papa and iss,suggesting it is an EHEC carrying multiple virulence genes.
The SYW004 strain showed high pathogenicity in ducklings.Experimental model of infection closely resembled the natural infection.Autopsy,histopathologic and ultra-structural pathological observation showed that the major pathological features were characterized by wide fibrinous inflammation in the serosa.Hyperemia,hemorrhage and inflammatory cell infiltrate were observed in heart,liver,spleen,lung,kidney,bursa of Fabricius,small intestine,pancreas and brain.Degeneration,necrosis or apoptosis were found in renal tubular epithelial cells,liver cells,cardiac muscle cells and intestinal epithelial cells.Lymphocytes decreased in bursa of Fabriscius.E.coli was observed on ultra-thin sections of heart,lung,spleen,liver and small intestine under electron microscope.The target cells were lymphocytes,phagocytes,intestinal epithelial cells and cardiac fiber.
6.Infection pattern of E.coli O46 isolate in an experimental model in ducks
To investigate the distribution of the O46 strain isolated in this study,an immunohisto chemical assay was developed with O46-specific antiserum and used to detect the bacteria of different organs at different times after experimental infection with the O46 isolate.The results showed this assay have good specificity.Samples of liver,spleen and kidney from ducks infected with O46 isolate showed positive results by this assay,while liver samples from dead ducks infected with E.coli O78,duck riemenella nantepestifer,duck plague virus,duck-origin avian influenza virus,salmonella or pasteurella multocida showed negative.With the advantages of direct,sensitivity and specificity,this assay could be used for accurate antigen localization.The study also indicated that the O46 isolates have wide tropism while heart,lung,spleen,kidney and intestine were found to be the main target organs.The bacteria were detected first in heart while bacteria could not be detected in trachea.Those findings provided important information for the in vivo distribution and proliferation of the O46 strain and accumulated useful knowledge for pathogenesis of this isolate.
引文
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